Julio Campo

Pulses of soil phosphorus availability in a Mexican tropical dry forest: effects of seasonality and level of wetting

Abstract Intact cores from the upper soil profile and surface litter were collected at the peak of the dry season and during the rainy period in the tropical deciduous forest of the Chamela region, Jalisco, México, to (1) analyze upper soil phosphorus (P) movement and retention, (2) compare soil P dynamic pools (soluble, bicarbonate, and microbial) in dry and rainy seasons, and (3) determine the response of these P pools to wetting. Unperturbed litter-soil cores were treated in the laboratory with either 10 mm or 30 mm of simulated rain with carrier-free 32P and compared to a control (no water addition) to determine the fate and retention of added P. 31P concentrations and pools in most litter and soil fractions were higher in the dry than in the rainy season. Soluble P was 0.306 g/m2 and microbial P was 0.923 g/m2 in the dry season (litter plus soil) versus 0.041 (soluble) and 0.526 (microbial) g P/m2 in the rainy season. After water addition, rainy-season cores retained 99.9 and 94% of 32P in the 10- and 30-mm treatments, respectively. Dry-season samples retained 98.9 and 80% of inputs in the same treatments. Retention after wetting occurred mostly in soil (bicarbonate and microbial fractions). Simulated rainfall on rainy-season soils increased P immobilization. On the other hand, simulated rainfall on dry-season soils released P through mineralization. The P release represents between 46 and 99% of the annual litterfall return. Our results suggest that both soluble and microbial P constitute important sources for initiation of plant growth at the onset of the rainy season in tropical dry forest.

Phosphorus cycling in a Mexican tropical dry forest ecosystem

The study was conducted in five contiguous small watersheds (12–28 ha) gauged for long-term ecosystem research. Five 80 × 30 m plots were used for the study. We quantified inputs from the atmosphere, dissolved and particulate-bound losses, throughfall and litterfall fluxes, standing crop litter and soil available P pools. Mean P input and output for a six-year period was 0.16 and 0.06 kg⋅ha−1⋅yr−1, respectively. Phosphorus concentration increased as rainfall moved through the canopy. Annual P returns in litterfall (3.88 kg/ha) represented more than 90% of the total aboveground nutrient return to the forest floor. Phosphorus concentration in standing litter (0.08%) was lower than that in litterfall (0.11%). Phosphorus content in the litterfall was higher at Chamela than at other tropical dry forests. Mean residence time on the forest floor was 1.2 yr for P and 1.3 yr for organic matter. Together these results suggest that the forest at Chamela may not be limited by P availability and suggest a balance between P immobilization and uptake. Comparison of P losses in stream water with input rates from the atmosphere for the six-year period showed that inputs were higher than outputs. Balances calculated for a wet and a dry year indicated a small P accumulation in both years.

Effects of Nutrient Limitation on Aboveground Carbon Dynamics during Tropical Dry Forest Regeneration in Yucatán, Mexico

Tree growth (as diameter increment), litterfall production, and litter biomass were studied in two secondary tropical dry forests of the Yucatán Peninsula under four treatments of nutrient addition. The study’s objective was to assess how variations in the nutrient supply affect aboveground net primary production and carbon (C) accumulation on the floor of two forests in different stages of regeneration. The study included an area of young forest (10 years old) with phosphorus (P)-poor soils and an area of old forest (around 60 years old) where soil P was comparatively less limiting. Four replicate plots (12 × 12 m) at each forest were either left intact (controls) or fertilized with nitrogen (N), P, or N plus P during 3 consecutive years. After 3 years of fertilization, relaxation of the constraints on nutrient limitation resulted in increased trunk growth rates at both the young and old forests. This effect was more pronounced with the addition of P or N plus P (trunk growth doubled with respect to controls), whereas N addition increased tree growth by 60% in comparison to trees in plots without nutrient supplements. In both forests, there were no significant differences in litterfall production among treatments during the first 2 years after fertilization. In the 3rd year of nutrient addition, litterfall production was significantly higher in plots fertilized with N plus P compared to control plots at both forest sites; however, changes in litterfall were not accompanied by litter accumulation in the floor of the two forests. The results of this study support the hypothesis that there is nutrient limitation during tropical dry forest regeneration. They further show that it may be maintained in the long term during secondary succession.

Leaf quality and herbivory responses to soil nutrient addition in secondary tropical dry forests of Yucatán, Mexico

Leaf quality (nutrient concentration) and herbivory were studied in a set of dominant tree species in regenerating, secondary tropical dry forests growing on limestone in the Yucatan Peninsula. The studys objective was to assess how variation in nutrient availability affects leaf nitrogen and phosphorus concentrations and herbivory. Our study included an area of young forest (10 y old), with phosphorus-poor soils, and an area of old forest (∼60 y old), in which soil P was comparatively less limiting. The foliage of representative species of each site was studied, including Acacia gaumeri , Leucaena leucocephala and Lysiloma latisiliquum at the young forest site, and A. gaumeri , Bursera simaruba and Pithecellobium dulce at the old forest. Four independent plots (12 × 12 m) at each forest were either left intact (controls) or fertilized with N, with P, or with N plus P for three consecutive years. Relaxation of nutrient limitations at the young forest resulted in an increase in leaf phosphorus and herbivory in all species. In contrast, N and P fertilization at the old forest site did not consistently affect leaf nutrient concentration and leaf damage. In this site the main response to fertilization was an increase in leaf nitrogen only in B. simaruba and an increase in herbivory in A. gaumeri ; leaf phosphorus was unaffected by soil fertilization. Overall, we observed that, subsequent to nutrient addition, leaf phosphorus concentration and herbivory in the leguminous species increased at the young site; leaf nitrogen concentration in the non-leguminous species increased at the old site while herbivory only increased in A. gaumeri , following a pattern similar to that observed in the young site. We conclude that the regulatory mechanisms of leaf quality and damage by herbivores will vary, depending on the details of the sites nutrient limitations and the identity of species.

Effects of nitrogen and phosphorus fertilization on the survival and recruitment of seedlings of dominant tree species in two abandoned tropical dry forests in Yucatan, Mexico

Seedling dynamics were studied in a set of dominant tree species in regenerating secondary tropical dry forests (TDF) growing on limestone in the Yucatan Peninsula. The objective of the study was to assess how variation in nutrient availability affects the recruitment and survival of individual species considering natural variations in light and bulk density in the topsoil. Our study included an area of young forest (10 years old), with phosphorus-poor soils, and an area of old forest (ffi60 years old). We used 16 plots (12 m � 12 m) per forest, in which we fertilized four plot replicates per treatment with nitrogen (N), phosphorus (P) and nitrogen plus phosphorus (NP). Another four replicates were kept as control. In four 1 m � 1 m sub-plots set of each experimental plot, tree seedling dynamics were studied over 2 years. The species with the highest recruitment were Acacia gaumeri and Leucaena leucocephala in the young forest, and Bursera simaruba and Phyllostyllon brasiliense in the old forest. Recruitment of A. gaumeri was affected more by light availability than fertilization, but its survival increased with N or with NP additions. Leucaena leucocephala benefited by fertilization (N, P, or NP) and the species regenerated on sites with lower light availability, but required high light conditions and N addition to increase its survival. The recruitment and survival of B. simaruba required low light availability, and the fertilization with P or with NP increased its regeneration. Phyllostyllon brasiliense was recruited only in the old forest. The addition of P resulted in the highest recruitment and the lowest survival among fertilization treatments, leading to a negative final seedling density. Canonical analysis revealed that in the young forest, the most important variables that influenced species recruitment were bulk density in the topsoil, fertilization with P, fertilization with N, and light availability. In the old forest, the P and N fertilizations and bulk density in the topsoil appeared to be the most important variables; only light was not significant. Light requirement differences between forests suggest that a large number of opportunistic species in the young forest could exist there. Overall, our results suggest that the dynamic of understory vegetation in Yucatan Peninsula is strongly influenced by nutrient availability, especially P. The responses of seedling dominant species to nutrient addition will vary, depending on the details of the sites light availability and soil bulk density, and the identity of species.

Tree seedling dynamics in two abandoned tropical dry forests of differing successional status in Yucatán, Mexico: a field experiment with N and P fertilization

The effects of nitrogen (N) and phosphorus (P) fertilization on seedling survival, recruitment, species richness and diversity in two abandoned tropical dry forests (10-yr old, young forest, and c. 60-yr old, old forest) in Yucatán, Mexico, were studied over two years. The seedling dynamics in the control plots were found to be highly seasonal with highest recruitment and lowest death rates during the rainy season. A low percentage of seedlings were resprouts; this important mechanism for forest regeneration had a higher-than-expected survival when compared to seedlings regenerated from seeds. Nutrient addition had significant effects on seedling dynamics in both of the forest regeneration stages. In the young forest, N fertilization facilitated the increase of seedling density. In the old forest, the addition of P decreased seedling diversity, while it increased the recruitment of only a few species. In both forests, P fertilization increased the survival time of seedlings when interacting with light availability and bulk density on the topsoil layer. Results suggest that low nutrient availability combined with low light availability constrain forest succession in Yucatán, Mexico.

Calcium, potassium, and magnesium cycling in a Mexican tropical dry forest ecosystem

We estimated the fluxes, inputs and outputs of Ca, K,and Mg in a Mexican tropical dry forest. The studywas conducted in five contiguous small watersheds(12–28 ha) gauged for long-term ecosystem research. A total of five 80 × 30 m plots were used for thestudy. We quantified inputs from the atmosphere,dissolved and particulate-bound losses, throughfalland litterfall fluxes, and standing crop litter pools. Mean cation inputs for a six-year period were 3.03 kg/ha for Ca, 1.31 kg/ha for K, and 0.80 kg/ha for Mg. Mean outputs in runoff were 5.24, 2.83, and 1.79 kg/ha, respectively. Calcium, K, and Mgconcentrations increased as rainfall moved through thecanopy. Annual Ca return in the litterfall (11.4 g/m2) was much higher than K (2.3 g/m2)and Mg (1.6 g/m2). Litterfall represented 99%of the Ca, 84% of the Mg, and 53% of the K, totalaboveground return to the soil. Calcium concentrationin standing litter (3.87%) was much higher than K(0.38%) and Mg (0.37%). These concentrations werehigher (Ca), lower (K), or similar (Mg) to those inlitterfall. Residence times on the forest floor were0.86, 1.17, and 1.77 yr for K, Mg, and Carespectively. Compared to the residence time fororganic matter at the site (1.31 yr), these resultssuggest slow mineralization for Ca in this ecosystem. Budget estimates were calculated for a wet and a dryyear. Results indicated that nutrients accumulated inthe dry but that nutrients were lost during the wetyear. Comparison of Ca, K, and Mg losses in streamwater with the input rates from the atmosphere for thesix-year period show that inputs are lower thanoutputs in the Chamela tropical dry forestecosystem.

Foliar nitrogen and phosphorus resorption and decomposition in the nitrogen-fixing tree Lysiloma microphyllum in primary and secondary seasonally tropical dry forests in Mexico

The tree Lysiloma microphyllum (Fabaceae) dominates in the seasonally tropical dry forests of central Mexico. In this study foliar N and P concentrations (on leaf mass basis), foliar N and P resorption efficiency and proficiency, as well as the decomposition of senescent leaves of L. microphyllum were studied in primary and in regenerating, secondary seasonally tropical dry forests. Our study included an area of early successional forest (10 y old), with phosphorus-poor soils and comparatively abundant nitrogen, an area of late-successional forest (∼60 y old), in which soil P and N were comparatively abundant, and an area of primary forest, in which soil P was comparatively abundant and N was less abundant than in the secondary counterparts. N and P concentrations in mature leaves varied across forests, reflecting soil nutrient availability. Nitrogen concentration in senescent leaves did not change among sites, which led to very different patterns of N resorption. In contrast, P concentration in senescent leaves was lower in the early than in late-successional and primary forests, which resulted in similar patterns of resorption. Leaf decomposition increased from 70% mass loss in the first year in the early successional to ∼80% in the same period in late-successional and primary forests. The element loss during decomposition change across forests in the following order: for N, early successional = late-successional > primary forest, and for P, primary forest > late-successional > early successional forest. Overall, the pattern of variation in leaf chemistry and nutrient release on the forest floor among sites is consistent with soil nutrient availability along this sequence, while decomposition rate may be related with the P concentration in senescent leaves.

Variations in soil carbon sequestration and their determinants along a precipitation gradient in seasonally dry tropical forest ecosystems

The effect of precipitation regime on the C cycle of tropical forests is poorly understood, despite the existence of models that suggest a drier climate may substantially alter the source-sink function of these ecosystems. Along a precipitation regime gradient containing 12 mature seasonally dry tropical forests growing under otherwise similar conditions (similar annual temperature, rainfall seasonality, and geological substrate), we analyzed the influence of variation in annual precipitation (1240 to 642 mm) and duration of seasonal drought on soil C. We investigated litterfall, decomposition in the forest floor, and C storage in the mineral soil, and analyzed the dependence of these processes and pools on precipitation. Litterfall decreased slightly - about 10% - from stands with 1240 mm yr(-1) to those with 642 mm yr(-1), while the decomposition decreased by 56%. Reduced precipitation strongly affected C storage and basal respiration in the mineral soil. Higher soil C storage at the drier sites was also related to the higher chemical recalcitrance of litter (fine roots and forest floor) and the presence of charcoal across sites, suggesting an important indirect influence of climate on C sequestration. Basal respiration was controlled by the amount of recalcitrant organic matter in the mineral soil. We conclude that in these forest ecosystems, the long-term consequences of decreased precipitation would be an increase in organic layer and mineral soil C storage, mainly due to lower decomposition and higher chemical recalcitrance of organic matter, resulting from changes in litter composition and, likely also, wildfire patterns. This could turn these seasonally dry tropical forests into significant soil C sinks under the predicted longer drought periods if primary productivity is maintained.

Leaf and litter nitrogen and phosphorus in three forests with low P supply

We compared the N and P contents of the main labile components of nutrient cycles in three different forest ecosystems [a tropical evergreen forest (TEF); a tropical dry forest (TDF); and a Mediterranean temperate forest (MTF)] with low P supply. A mass-balance approach was used to estimate mean residence times for organic matter, N and P in the forest floor, and to examine the flexibility of N and P intra-system cycling in the three forest ecosystems. For this purpose, we combined published values of N and P in foliage, litterfall, forest floor litter and mineral soils in these three forest ecosystems. The results of our analysis were consistent with the widely held belief that the N content of leaves (both green and senescent) and litter increases with increasing temperatures. In contrast, the data did not support the hypothesis that leaf P content decreases with increasing temperatures and precipitation: leaf and litterfall P contents were higher in both tropical forests than they were in the temperate forest. The TEF had the highest P content of the three forests studied. The mass-balance analysis indicated that although P mineralization in the TDF can run ahead of litter decomposition stoichiometry when P is in short supply, flexibility is much reduced or absent in the TEF and the MTF. Our analysis provides additional evidence of the importance of climatic factors in forest ecosystem processes and highlights the role of flexibility in ecosystem nutrient cycling, especially for P in ecosystems with a limited P supply.